Vision



Vision





Like hearing, vision depends on our being able to analyse vibrations: not of mechanical waves, but of electromagnetic ones. This implies an entirely different kind of transduction mechanism, and – as always – some understanding of the underlying physics.




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Light and dark

Light is a form of energy propagated by electromagnetic waves travelling at an immense velocity – some 300 km/ms – and carried in discrete packets called quanta or photons. As you can see above, only a very small range of all the wavelengths of electromagnetic radiation is visible. The longest waves that we can just see, forming the red end of the spectrum, are some 0.7 µm in length, slightly less than twice as long as the shortest waves at the blue end. In nature, most electromagnetic radiation is generated by hot objects: the hotter they are, the more of this energy is radiated at shorter wavelengths. The peak of the spectrum of light from the sun – an exceedingly hot object – corresponds quite closely with the range of wavelengths seen by the cone receptors in the eye (the expanded graph above). Of man-made sources of light, many, like the ordinary incandescent electric lamp, radiate as hot bodies and have a smooth and broad emission spectrum; others are quite different, and emit light only at a few discrete wavelengths. The older sodium lights used for street lighting, for example, are effectively monochromatic, their energy being concentrated in a very narrow band in the yellow region. Domestic fluorescent lamps have a spectrum consisting of a number of emission lines superimposed on a continuous background.

The spectrum in a sense defines the quality of a light; determining its quantity is called photometry, and is complicated by the fact that there are two kinds of photometric measurements: first, how much light is emitted by a source of radiation (which may be selfluminous, like the sun, or illuminated, like the moon), and second, how much light is received by an illuminated object. The candela (cd) is a measure of the rate of emission
of light by an object: an ordinary 60 W bulb emits about 100 cd. The amount of light received by an object per unit area is its illuminance, and is measured in lux. This unit is defined as the degree of illumination of a surface 1 m from a source of one candela radiating uniformly in all directions. Full sunlight may provide about 100 000 lux.




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Now objects in the real world scatter back some of the light that falls on them, so that in general an illuminated surface is also a luminous one, emitting a certain amount of light per unit area: this is described by its luminance, measured in candelas per square metre (cd/m2). Finally, the ratio of luminance to illuminance in these conditions is a measure of the surface’s whiteness or albedo. If we shine one lux on a perfectly white object that is also a perfect diffuser, it will have a luminance of about 0.32 cd/m2, and such a surface is said to have an albedo of unity. Ordinary white paper has an albedo of about 0.95; paper printed with black ink, about 0.05. The photometry of coloured objects, which scatter back light of a different spectral composition from that which illuminates them (so that their albedo is a function of wavelength) is much more complex and requires special definitions and methods of measurement. image




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This chart gives some idea of the range of luminances found in nature. At the bottom end, the eye can function at light levels measured in terms of single photons; at the upper end, the brightest lights we can tolerate without retinal damage are an amazing 1015 times – 15 log units – stronger than this. These are extremes: the receptors in the eye, the rods and cones, provide useful vision over the middle 1012 or so of all this. This is an extraordinary performance that no man-made device can begin to challenge: in television studios, for instance, absurdly high levels of lighting have to be used to produce decent pictures. So here is something very special about vision – a function whose technical name is light and dark adaptation or simply adaptation for short.


Adaptation: a sliding scale

In practice, however, at any one moment the actual range of luminances to which the eye is exposed is very much smaller than this. The albedos of natural objects vary only from about 0.05 to 0.95. As you look round a uniformly illuminated room, the range of luminances that you see is therefore only about 20:1. This is true whatever the illumination: whether a room is bathed in sunlight, or dimly by light-bulbs, black objects still look black and white objects, white. Black and white are thus relative terms: the eye operates on a sliding scale of brightness that can be moved up and down the whole range in such a way as to match the prevailing level of luminance; this property is the result of various mechanisms of adaptation.




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As a result, the range of the visual system is dynamically matched to the range of luminance in your visual surroundings. If the general illumination changes, the visual system quickly follows suit. It follows that the eye responds not so much to the luminance of natural objects as to their albedo: a much more useful sensory quality, since albedo is an intrinsic property of objects that helps us to identify them, whereas their luminance depends
on how much they happen to be illuminated. A brightly illuminated patch of black on a white background still looks black, even though its luminance may be more than a neighbouring dimly illuminated patch of white. So adaptation has two functions: it enables you to cope with a very wide range of light levels, but it also provides information about albedo, the first step towards recognition.




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Contrast

The existence of adaptation has profound effects on what we are able to see and how things look. It is easy to show that in a very wide variety of situations our perceptions are scaled to the prevailing luminance level. Much of the time we are trying to see small differences of luminance on a relatively uniform background, and in many situations it turns out that their perception depends crucially on something called contrast. Contrast is the ratio ΔI/I between some difference in luminance ΔI and the background luminance I. It can be defined equally well for positive or negative increments on a steady background (a star in the evening sky, a fly on a sheet of paper), or for the repetitive stimuli often used by visual scientists called gratings in which the luminance varies – often sinusoidally – around a mean level I. In general, we detect an object if its contrast exceeds a certain threshold value, otherwise it will be invisible. This explains, for instance, why we may be able to see a very thin white line on a black background, yet not see a black line of identical width on a white background. Though ΔI is the same in each case (but of opposite sign), I is not, so the contrast is considerably bigger for the dark background. Or consider stars, which have a fixed ΔI: as the sun rises, I increases, their contrast drops, and one by one they disappear. In good conditions, threshold contrast is typically around 1 per cent.




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Photopic and scotopic vision

Several distinct mechanisms contribute to this ability of the eye to adapt to the prevailing level of illumination, and they are discussed in more detail on p. 147. Some respond quickly to a sudden change in the ambient level, others more slowly. If we go from daylight to a dark room we find that it takes nearly 40 minutes for the eye to adjust its sensitivity fully to the reduced level of illumination. The simplest way to demonstrate this process of dark adaptation is to plot a graph of a subject’s absolute threshold – the luminance of the dimmest light he can just perceive – at regular intervals during this adapting period. Such curves normally show two distinct components: an initial one that levels off after some 8 minutes, and a further, slower increase in sensitivity that takes another 30 minutes or so to reach completion. image




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The points in the graph above show absolute thresholds at different times after presenting a strong light that bleached 24 per cent of the pigment in the rods, and the coloured line shows the time-course of the recovery of sensitivity of the cones alone. So apart from demonstrating the slowness of dark adaptation, the two-stage recovery also demonstrates the existence of two populations of receptors in the retina.



  • Cones are found particularly in the middle of the visual field and provide very detailed information about the retinal image, being a little more than 2 µm in diameter, and are also responsive to colour. But cones have a high threshold, and can only function when the light is above some 10-2 cd/m2 (the photopic region). Below this, in the scotopic region, we’re forced to use the rods. At in-between levels we have an intermediate kind of vision called mesopic. The top of this mesopic region is at about 100 cd/m2, when the rods stop functioning because they’re completely saturated.


  • Rods are much more sensitive – in fact as sensitive as they could possibly be, since one individual rod can respond to a single photon of light – but the bad news is that in order to achieve this sensitivity they have to group themselves together into functional teams – numbered in thousands – by means of their neural connections in the retina. By pooling their information they enormously increase their sensitivity (whereas the random background noise tends to cancel out by being pooled, the signal one is trying to detect does not). But this is at the cost of throwing away a lot of information about the spatial detail of the retinal image, and also sacrificing the ability to distinguish wavelengths or colours; they are also slow, and they respond to a range of wavelengths that is slightly shifted in the blue direction – something called the Purkinje shift. image The difference in the spectral sensitivity curves relates to the degree to which rods and cones absorb light of different wavelengths: the dashed green line shows how the absorption spectrum for the pigment in the rods, rhodopsin, has a very similar shape to the scotopic sensitivity.








Table 7.1 Vision under photopic and scotopic conditions
































Photopic


Scotopic


Sensitivity


Low; best vision in fovea


High; best vision outside fovea



Light entering periphery of pupil less effective than centre (Stiles-Crawford effect)


No Stiles-Crawford effect


Spatial properties


High acuity; contrast sensitivity reduced at low spatial frequencies (lateral inhibition)


Low acuity; less lateral inhibition


Temporal properties


High flicker fusion frequency; reduced sensitivity at low frequencies (fast adaptation)


Low flicker fusion frequency; less fast adaptation. Increased latency


Wavelength


Most sensitive at around 550 nm


Most sensitive at around 500 nm (Purkinje shift)



Trichromatic colour discrimination


Monochromatic





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So we have a sort of dichotomy between these two types of vision. Some of these differences between photopic and scotopic vision are summarized in Table 7.1 and further explanations of the terms used are given in the sections that follow.


Image-forming by the eye


Optics

The eye is not just a device for sensing light and dark: it forms an image of the outside world, and encodes it as a spatial pattern of neural messages for the brain. Image formation is a matter of basic physics: when light enters a region of higher refractive index (or leaves one of lower) it is bent towards the normal, by an amount that depends on the refractive index, µ. So if the surface is curved, the further out bits bend parallel rays more and the inner
ones less, and if you’re lucky they come to a point. To a first approximation, the way to get them to come to a point is to use a spherical surface. In the eye, there are three surfaces of this sort that act together to bring the images of distant objects to a focus on the retina: they are the cornea, and the front and back surfaces of the lens. The refractive index of the aqueous humour that separates the cornea and lens is much the same as that of the vitreous humour that fills the rest of the eye, and is about 1.34; that of the crystalline lens is only slightly greater than this, being about 1.41.




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Opticians describe the power of refractive surfaces by the reciprocal of their focal length in metres, and these units are called dioptres (D); for a concave lens, the power is negative. An advantage of dioptres is that they add up: if you have several lenses in series, the total power is simply the sum of the powers of each individual component. Since the distance from the cornea to the retina in Man is about 24 mm, you might think the total refractive power of the eye when focused on a distant object would be some 43 D; but because the focal length is being measured within a refractive medium, this figure must be multiplied by the refractive index (1.34), giving about 57 D. Of this, about 48 D are due to the cornea alone, and only 19 D or less to the lens. image So, contrary to popular belief, the focusing power of the eye is not mainly due to the lens: the reason is that its refractive index is not very different from that of the aqueous humour and vitreous humour on each side, so it doesn’t contribute much. The cornea is powerful because its interface is with air, whose refractive index is much smaller (1.0008 to be exact).




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Accommodation and the lens

Though the lens contributes little to the total refractive power, it still has a very important function: it can alter its shape and hence fine-tune the eye’s effective focal length; this function is called accommodation. It’s able to do this because it is elastic and flexible, made of remarkably long (12 mm) thread-like cells, rectangular in cross-section and – for transparency – lacking nuclei, knitted together in a series of concentric layers by zip-fastener-like balland-socket joints that provide flexibility. You can think of the lens as a kind of jelly held in place by guy-ropes all round the edge, the suspensory ligaments that are normally under tension from elastic elements attached to the wall of the eye and tend to flatten the lens out. Encircling the lens is a ring of muscle called the ciliary muscle. When it contracts, it relieves the tension on the filaments, and the lens relaxes into a more bulging, rounded shape with a shorter focal length.




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The lens presents something of design problem for
nature, since – being transparent – it obviously cannot have a blood supply. It obtains its nutrients and oxygen from the aqueous humour that bathes it on both sides, and is able to penetrate the lens because of fibrous nature. It is a fluid similar to plasma but with only some 1 per cent of its protein concentration and a peculiarly large amount of ascorbic acid. The aqueous humour is continuously secreted by the ciliary body, and passes through the iris into the anterior chamber where it eventually filters its way out into the canal of Schlemm, whence it ends up in the bloodstream. The resistance to its outflow generates an intraocular pressure of some 10-20 mmHg. Blockage may raise this pressure to the point where the flow of blood into the back of the eye is hindered, a serious condition called glaucoma which is a common cause of blindness.




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The range of accommodation is the difference between the maximum and minimum power of the optics, expressed in dioptres. It can easily be measured by finding the positions of the near and far points of the eye, the nearest and furthest distances at which objects can just be brought into focus. For a normal or emmetropic eye with accommodation fully relaxed the far point will be at infinity (0 D), and the range of accommodation will be given by the reciprocal of the distance of the near point in metres. For instance, a young emmetrope’s near point will generally lie at around 80 mm, so that the range of accommodation is 12 D. As one gets older, however, the lens begins to seize up, so that it no longer bulges so much when the ciliary muscle contracts, and the range of accommodation falls. By the age of 60 the possible amplitude of accommodation may have fallen to 1 D or so, a condition known as presbyopia. It is the near point that moves further away, and when you find you can no longer read the newspaper even at arms length, then you have to start wearing either half-glasses, or bifocals, in which the bottom half of the lens has a stronger power than the top. image




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Errors of image formation

That’s the basic optics. How good are they? image Few people are exactly emmetropic; for most, one finds that when the accommodation is fully relaxed the total refractive power is either too strong or too weak in relation to the distance from the cornea to the retina: they suffer from refractive error. If it is too strong, the image of a distant object lies inside the vitreous instead of on the retina and we have the condition called myopia. image An optician corrects myopia by using a negative or concave spherical lens. For example, if your far point is 1 m away, then that means your eye has 1 D too much power, and you need a -1 D lens to make it up. If you’re long-sighted or hypermetropic, the eye is too weak and needs an additional positive lens to correct it. In either case, you can describe the degree of disability by the power and sign of the lens needed to bring the eye back to emmetropia: thus a mildly short-sighted patient might require a correction of -1.75 D. This is called the spherical correction, part of an optician’s prescription, and in general is not the same in both eyes.




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Astigmatism

A more subtle but very common type of focusing error occurs when the curvature of the cornea is different in different meridians. It will then focus a horizontal line at a different focal length from a vertical one, a condition
called astigmatism. If, for example, it has a smaller radius of curvature in the horizontal plane than in the vertical, the far point when measured with a vertical line as test object will be closer than when a horizontal line is used. It is corrected with a cylindrical lens – in effect a section cut from a cylinder, just as a spherical lens is from a sphere: it focuses only in one meridian. Opticians test for astigmatism by means of a target like the one below, called an astigmatic fan; an astigmatic subject will see some of the lines more sharply than others, and this will tell the optician the angle at which a cylindrical lens should be placed in front of the eye to make the refractive power as nearly as possible equal in all meridians. The power of the cylindrical lens that is needed to do this, together with its meridional angle, make up the cylindrical correction that is the second part of a prescription for spectacles. image




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Astigmatism and incorrect refractive power are not the only faults that may be found in the eye’s optics, and – as in many man-made optical systems – the cornea and lens together produce a number of different types of optical defects in addition to refractive error.


Chromatic aberration

The first of these is due to the fact that the refractive indices of the various optical media of the eye depend on the wavelength of the incident light. In general, the refractive index increases with decreasing wavelength, so that blue light is refracted more than red. This phenomenon, called dispersion, means that the focal length of a lens depends on the wavelength, with blue being shorter than red, amounting to some 2 D over the whole visible spectrum. This gives rise to defects in the resultant image, in the form of coloured fringes, called chromatic aberration. So if you look at a blue object and a red object lying side by side at the same distance from the eye, they cannot both be in focus simultaneously, and a subject who is emmetropic when his far point is measured in red light will be short-sighted if it is measured in blue: his far point will then be only a metre or so away. This forms the basis of a simple clinical test for errors of refraction, consisting of an illuminated screen divided into three portions that are red, green and white: identical test figures are superimposed on each field, and the subject is simply asked which figure he sees most clearly. A myope, whose lens is too strong anyway, sees red targets more clearly than green, and vice versa for a hypermetrope; an emmetrope will see the one on the white background best.




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The importance of chromatic aberration can be seen by the fact that visual acuity is improved by some 25 per cent in monochromatic yellow light. The eye mitigates the effects of chromatic aberration in two ways: by a yellow pigment over the fovea that reduces the blue component, and by the fact that very few blue cones are found in the centre of the fovea, where the finest spatial vision is found.


Spherical aberration

The other aberration is spherical aberration. We saw earlier that to make a surface bring parallel rays to a point it needs to be roughly spherical. Although ordinary
man-made lenses are nearly all spherical, simply because they’re easier to make, a spherical lens does not in fact bring rays to a point focus: they get bent too much as you go further out, and it is the resultant blur that is spherical aberration. The shape you really need is not a sphere but an ellipsoid. For surfaces that are small in comparison with their radii of curvature the difference is slight, and spherical aberrations are often negligible. But in the case of the eye, the aperture is of the same order of magnitude as the radius of curvature of the cornea, and the result is that rays entering near the periphery of the cornea are bent too much, and form a closer focus than those entering near the centre. To some extent Nature has compensated for spherical aberration, first of all by making a cornea that is not exactly spherical but tends towards the desired ellipsoid; and second, in that the refractive index of the lens is not constant throughout, but graded from a maximum of some 1.42 at its centre to about 1.39 at the edge, thus cancelling out, to some extent, the extra bending of peripheral light rays. The degrading effects of both spherical and chromatic aberration, and of other defects due to irregularities of the refracting surfaces, get worse as the pupil or aperture of the eye increases, and this in turn depends on the state of the iris: this is discussed on p. 137, in the context of visual acuity. image




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Scatter and diffraction

There are two other problems with the eye’s optics that are not exactly errors of focus, but do degrade the quality of the retinal image. When light enters the eye it tends to get scattered, particularly by the cornea and lens but also to some extent by bouncing off the back of the retina. This effectively superimposes on the image a more or less uniform background, perceived as glare, whose illuminance is of the order of 10 per cent of the mean illuminance of the retina. This means that if we look at a target whose actual contrast is 100 per cent, the effect of this scatter is to reduce the contrast of the retinal image to something nearer 90 per cent. Because of the progressive opacity of the lens, glare gets worse as you get older; it is particularly obvious when driving at night in the face of opposing headlights. The second problem is something that is caused by the pupil, called diffraction. Whenever light passes through a restricted aperture it tends to spread out and therefore degrades the retinal image: the smaller the aperture, the worse this gets. More precisely, the width of the resultant pointspread function is of the order of λ/d radians, where λ is the wavelength, and d the aperture of the system. In practice, so long as the pupil is bigger than some 3 mm, diffraction contributes rather little in comparison with the other optical problems.


The pupil

Thus the ideal size for the pupil is a compromise. An important factor is the ambient light level: under bright photopic conditions the eye can take advantage of the excess light by constricting the pupil and improving the quality of the retinal image. In scotopic conditions, however, the eye needs all the light it can get and the quality of the retinal image is of secondary importance: in any case, we shall see later that the rods are not capable of passing on accurate information about the detailed structure of the retinal image. It is important to emphasize that pupil dilatation contributes very little to the enormous changes in sensitivity that accompany dark adaptation, since it can only vary the incoming light by a factor of 16 at most, or 1.2 log units. The effect of pupil diameter on visual acuity is discussed in more detail on p. 137.

Another factor is that the control of the pupil is closely linked to accommodation: when the ciliary muscle contracts in order to focus on a near object, there is normally an associated constriction of the pupil (the near reflex), which may help to increase depth of focus (see p. 137). As these responses are usually also combined with binocular convergence movements of the two eyes, the whole pattern of response (constriction, accommodation, convergence) is also known as the triple response. Under certain clinical conditions, notably in neurosyphilis, one may find that the pupillary response to near objects remains despite loss of the response to bright lights: this condition is known as the Argyll Robertson pupil, and is an important diagnostic neurological sign. The fact that pupil dilation is also a measure of general sympathetic activity and of emotional or sexual excitement also has it uses. image


Visual acuity


Measurement

Visual acuity is a measure of the fidelity with which the visual system can transmit fine details of the visual world: it is the equivalent of the ability of a camera to produce sharp pictures. In a camera there are essentially two stages at which sharpness may be lost: either through optical defects that blur the patterns of light in the image, or subsequent factors – a lack of pixels – that limit the density of detail. These correspond in the eye to the quality of the optics, and to the density of the retinal receptors. But in the case of the eye there is a third factor: the possible degradation of the image that may occur in the course of the neural processing that takes place in the retina.




The pointspread function

The effect of optical blur is relatively straightforward. Consider for instance the simplest of all visual objects, a star. Stars are so far away that they can in effect be regarded as infinitely small point sources. But the retinal image of the star will certainly not be a point, because the optics will spread the light out into a sort of heap on the retina. This distribution of light is called the pointspread function, and its size is a useful measure of how good or bad the optics are. In the human eye, under the best possible conditions,
the pointspread function has a diameter of about 1.5 arc min (measured half-way up); the worse your optics, the bigger this becomes. image




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This pointspread function is of fundamental importance, since it absolutely determines what sort of patterns we can see and what we can’t. For instance, if we have two stars rather than one, each with its own pointspread, then if they are far apart, they will be seen correctly as two separate stars; if closer, eventually there will be no little dip in between them, and the brain will have no way of knowing that there are two stars and not one. For normal observers, the angle of separation for which this kind of resolution can just be performed provides a quantitative measure of visual acuity. Its value – around 30-45 arc sec – is an order of magnitude greater than the width of a black line that can just be seen. This figure, that is found to apply in many similar tasks of resolution, can be taken as a measure of the visual acuity or resolving power of the eye.




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Resolution

We need to pause at this point to consider exactly what is meant by ‘seeing’ something. Seeing can mean detection, or resolution, or recognition. An ornithological friend points up in the sky and says ‘Can you see the crested willow-warbler?’ If you can’t, it may be either because it’s so far away you can’t detect it, or because although you can see a formless speck you can’t resolve the pattern of its markings, or finally because – although you can perceive every aspect of it – you haven’t the least idea what a crested willow- warbler is meant to look like. In each case, there is a failure to ‘see’, but for completely different reasons.

The existence of spatial spread has implications for detection as well as resolution. Since the incident energy from the point source is spread out over a larger area, the maximum intensity at the central peak is necessarily reduced, leading to a decrease in the contrast, ΔI/I, which determines whether it will be seen against its background (see p. 128). For objects of intrinsically high contrast, such as stars seen against the void of space, this will not matter much, and subjects with poor visual acuity as measured conventionally (see below) are not as bad at seeing stars as one might expect, bearing in mind the fact that such objects subtend an almost infinitely small angle. In the dark, whether one sees a star or not is almost entirely a matter of whether a sufficient number of photons from it fall upon a rod summation pool; as the sun rises its visibility depends on whether ΔI/I exceeds the threshold contrast. Thus – rather as in the case of the skin, discussed on p. 90 – we may find that we can localize a visual object to a much higher degree than our ability to tell whether there is one object or two.

While the main effect of contrast is on detection, it also has some effect on resolution. With the two stars, in a case like that of (c), it is clear that we cannot improve resolution simply by increasing the contrast, and such a stimulus may be described as absolutely unresolvable. But in an intermediate case like (b), whether resolution is possible or not will depend on the contrast of the original object as well as on the width of the pointspread function. This interaction between resolution and contrast can best be investigated by using grating patterns as test targets. A grating is simply a regular pattern of stripes; if the stripes are uniformly black and white, it is called a square-wave grating, because a plot of intensity as a function of distance across the grating would have a square-wave profile. In the same way, sinusoidal gratings have an intensity profile that is sinusoidal: there was an example on p. 128. In each case, one can describe the grating in terms of its spatial frequency (i.e. the number of cycles per degree) and its contrast (defined as the difference in intensity between peak and mean intensity divided by the mean intensity). Thus a pattern of alternate pure black and pure white strips, each 1° across, could be described as a square-wave grating of 100 per cent contrast and spatial frequency 0.5 cycles per degree. A simple experiment is to ask a subject to view a sinusoidal grating of a particular spatial frequency, and then reduce its contrast until he reports that he can no longer see it. If we plot this threshold contrast as a function of spatial frequency, we typically obtain a curve such as the one below. Because a blurred pointspread function affects high spatial frequencies much more than low ones, the contrast required to see the grating increases sharply as its frequency is increased, until at about 40-50 cycles per degree (the cut-off frequency), the subject cannot even see a grating of 100 per cent contrast. Because of the steepness of this cut-off, a small amount of extra blur causes a large increase in the
contrast needed, and so the method provides a sensitive measure of acuity. The reason for the fall-off in contrast sensitivity at low frequencies is discussed later (p. 152).




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Of course, dispensing opticians don’t bother with all that. A rough-and-ready measure of visual acuity is to make up charts of letters of standardized shape and graded in size, and see at what point the patient is unable to read them. A common chart of this kind is the Snellen chart, in which rows of letters of diminishing size are to be read: by discovering the row at which the subject finally stops. Knowing the size of letters and the subject’s viewing distance, one can estimate his minimum resolvable angle. You can see that a letter E, for instance, is a bit like a miniature grating, and you need to be able to resolve it in order to read it. Each line of text is marked with distance in feet at which you should just be able to read it. If at 6 m you can only read the line for 12 m, then your acuity can be described as 6/12; a normal person is in theory therefore 6/6. image But Snellen charts are very conservative, since they are calculated not on the basis of 45 but of 60 arc sec, or one minute. So at 6 m away, with very good vision you should in fact be able to read the line marked 5, in which case your vision is described as 6/5. The idea is no doubt to make the glasses the optician has just sold you seem better than they are.

The difficulty of the Snellen chart for scientific work is that the test is only partly one of resolution. Apart from the assumption that you are literate, it is also clear that some letters are recognized more easily than others because of their overall shape; and for some purposes the Landolt C chart, used in the same way, is preferable because it provides no extraneous clues to the subject. Even this is not ideal, since it is still possible to detect the overall orientation of the C even though it is not really resolved: for this purpose, simple barred patterns are better, since when blurred it is completely impossible to guess the original pattern, which is not true either of Snellen letters, or even Landolt Cs. Another kind of chart in which all the letters are the same size but are graded in contrast can be used to test for certain kinds of defects in the visual system in which sensitivity to contrast is specifically impaired.




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The tests described so far are all genuine tests of acuity in that they require that detail of some kind be resolved. Other tests, that at first sight might also appear to be acuity tests, are really tests of detection or localization, and give apparent acuities far better than 30-45 arc sec. A well-known example is vernier acuity, where a subject is required to move two lines into alignment, as for instance in the scale of vernier callipers. Here one does incredibly well, typically of the order of a few seconds of arc. But the task is not resolution but localization: even if
the retinal image is blurred, one can still estimate where the peak is quite accurately. One can show that the longer the line the better one is, showing that accuracy is also being improved by averaging information over the whole line. Another pseudo-acuity task is the detection of stars, which may subtend extremely small angles at the eye. For instance, the bright star in Orion called Betelgeuse subtends only some 1/20 arc sec. But in a sense that figure is quite irrelevant: because of the pointspread, its image still has a width of 45 arc sec, and whether you detect it or not depends simply on how whether its luminance exceeds the absolute threshold, ΔI0. image




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The influence of the pupil

Unlike the lens, the size of the pupil is under the control of two different muscles: one, the sphincter pupillae, lies circumferentially round the iris, and the other, the dilator, lies radially. So the two muscles have opposed effects, the first causing contraction of the pupil and the second dilatation, and they are respectively under the control of the parasympathetic (grey in the diagram below) and sympathetic systems (blue).

It is not entirely clear which of the two branches of the autonomic nervous system is responsible for normal tonic control of pupil size, and one may cause mydriasis (enlargement of the pupil) by drugs that either block the action of acetylcholine on the sphincter (e.g. atropine) or simulate the effect of noradrenaline on the dilator (e.g. phenylephrine). Light causes constriction through the parasympathetic route from the ciliary ganglion, in turn activated by the Edinger-Westphal nucleus high up in the brainstem, close to the oculomotor nucleus, which receives sensory information about overall light level from neurons in the pretectum, which themselves receive fibres from the optic nerve.




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The advantages of a large pupil size are first that the eye receives more light (over the normal range of pupil diameters, about 2-8 mm, the amount of light caught by the eye varies by a factor of 16), and second that the diffraction effects that always occur when light passes through a small aperture are minimized. The advantages of a small pupil, on the other hand, are an increased depth of field (a greater tolerance of errors of focus, as can be seen in the diagram) and a reduction in the magnitude of the optical aberrations and of glare: the extent of this effect can be seen for oneself by looking through a pinhole, which acts as a very small artificial pupil. Aberrations are reduced because the smaller the pupil, the more nearly the optical surfaces will approximate to their ideal forms, and the less noticeable the aberrations will be. The effects of diffraction can be calculated without much difficulty. For a pupil of diameter 2.5 mm, and with green light, diffraction alone creates a pointspread of a little less than one minute of arc. In other words, under these conditions acuity is effectively limited by diffraction at the pupil. In the dark, with a pupil of some 8 mm diameter, the corresponding figure for diffraction alone is about 17 arc sec, but the actual pointspread is very much wider than this because of the increased contribution of the aberrations when the lens is widely exposed: in fact the pointspread function actually gets wider with increasing pupil diameter beyond 3 mm or so. As a result, a graph of visual acuity as a function of pupil size is U-shaped, with a distinct optimum around 3 mm. Thus if acuity were the sole consideration, we might expect to find the pupil always fixed at that value. But as we shall see, under
conditions of dark adaptation the intrinsic acuity of the neural processing of the retinal image is so low that the poor optics contribute little to the overall blur, and the advantage of being able to increase retinal sensitivity by catching more light with a dilated pupil outweighs the disadvantage of slightly decreased acuity.




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Table 7.2 Advantages and disadvantages of small and large pupils














Smaller pupil


Larger pupil


Increases depth of field


Receives more light


Minimizes optical aberrations


Minimizes diffraction effects


Minimizes glare



The influence of the retina

The previous section may have given the impression that acuity is purely a matter of the optics, and that what the retina and brain do doesn’t matter very much. Under photopic conditions, for most of us, in practice this is probably true. But even when there is an abundance of light, and the optics are free of any defects, then one finds that the visual acuity that one measures is not quite as good as one would expect from the quality of the retinal image; what is happening is that the retina and brain are introducing some extra deterioration of their own.

The most obvious way in which the retina influences visual acuity is simply that the receptors are a finite distance apart from each other, which in itself inevitably limits the detail that can be seen. In the very centre of the fovea receptors are at their closest spacing a little over 2.3 µm, or half a minute of arc, which happens to be about the same as the half-width of the pointspread. Actually this is no coincidence, since obviously it wouldn’t make much sense to have wonderful optics with a beautifully detailed image combined with huge receptors incapable of transmitting the information; nor would it be helpful to have finely packed receptors able to respond to details that in real life they could never possibly experience, because of optical blur.




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The relative importance of optical as opposed to retinal and neural factors in determining acuity can be determined directly by arranging to project a grating on the retina in such a way that its contrast is unaffected by the quality of the optics. One way of doing this is to generate interference fringes on the retina by means of two point sources of coherent light from a laser: the resulting interference pattern is in effect a sinusoidal grating, whose frequency depends on the separation of the two sources, and whose contrast is substantially unaffected by the quality of the optics. One can then measure the subject’s threshold contrast as a function of frequency, as already described, and compare the result with what is found when viewing a ‘real’ sinusoidal grating. Although there is some improvement when the optics are bypassed in this way, even when the eye is fully corrected it is not a very great one. This suggests that the retina is in a sense matched to the eye’s optical properties.

But it’s not just the receptor spacing that counts; we need to consider what happens to the neural signal as it goes through the retina to the brain. In many ways one can think of the spatial pattern of activity in the receptors and subsequent neurons as another kind of image – a neural one – that may be subject to the same kinds of degradation as an optical image.


Neural blur

If the receptors simply had a one-to-one connection to bipolars, and bipolars to ganglion cells, then one would not expect any deterioration to occur as the neural image is passed along. We shall see later that this is perfectly true in the centre of the fovea, where acuity is highest, but is certainly not the case further out. Here one finds many receptors pooling their information, funnelling onto one bipolar, and many bipolars funnelling onto one ganglion cell. A telling statistic is that each eye has about 130 million receptors but only about one million ganglion cells. In fact the degree of convergence in the periphery is actually even bigger than that implies, because of huge overlap of receptive fields. In the periphery, a typical alpha ganglion cell may receive input from a staggering 75 000 rods, via 5000 rod bipolars. This is good news from the point of view of trying to detect things, because it increases the difference between the size of the signal you are trying to detect, and the background noise which would otherwise obscure it: in fact it accounts for nearly
all the difference in sensitivity between rods and cones (since the difference in threshold between one isolated rod and one cone is only about a log unit). But it is not so good for acuity. Because this neural pointspread is broader the further from the fovea, acuity is very much worse in the periphery than in the centre. And most important of all, as you dark-adapt, changing from central cone vision to rod vision with their enormous pooling of information, acuity drops off dramatically. If the contrast threshold as a function of spatial frequency is measured with a fixed pupil during progressive stages of dark adaptation, over 6 log units one finds a steady decrease in the cut-off frequency, the result of changes in the neural organization of the retina. One of the adaptational responses to reduced light levels, as we shall see, is an increase in the effective size of the ganglion cells’ summation pools, so that they can catch more light. But this obviously has the effect of increasing the degree of neural blur, and hence of reducing the overall acuity.




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And this brings us back to the pupil once again. The fact that as you dark-adapt, the neural blur introduced by the retina increases, and begins to dwarf the real optical blur caused by bad optics, means that the pupil can now afford to get bigger. Whereas a large pupil in bright light is a very bad thing because of the effect it has in increasing the aberrations and defects of focusing, in dim light these are not so important, so that one can enjoy the benefits of having more light for the purposes of detection. Thus there is a kind of necessary reciprocal relation between sensitivity and acuity; the better you are at one, the worse you are at the other. A big pupil provides more sensitivity but worse acuity; pooling of receptors onto ganglion cells also provides more sensitivity but less acuity. We shall see other examples of this kind of trade-off later on.

To summarize, there are many factors that contribute to visual acuity, and their relative contributions depend on the state of adaptation of the eye: they are summarized in Table 7.3. In good light, an emmetrope’s acuity is limited by diffraction, and thus about as good as could be expected from an eye of the size that we actually have. But most people are not emmetropes, so that without spectacles it is refractive error that limits their acuity.


The retina

One might hope to be able to see another person’s retina directly by eyeball-to-eyeball confrontation: if both eyes are emmetropic and relaxed, each retina should be clearly in focus on the other. The reason why this doesn’t in fact work is that the presence of the observer’s eye also prevents light falling on the other’s retina, so that nothing can be seen: under normal conditions the pupil of the eye is always dark. The ophthalmoscope is a device that gets round this problem by projecting a small beam of light from a light source (S) into the subject’s pupil at the same time. It also has an arrangement whereby one of a set of negative and positive lenses (L) can be introduced into the optical pathway: the power of the lens that exactly brings the subject’s retina into sharp focus is equal and opposite to the combined refractive errors of observer and subject. Thus so long as an oculist knows his own correction, the ophthalmoscope provides an objective method for determining what spectacles the subject requires, as well as permitting the examination of the retina for signs of disease.




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Two features of the retina are immediately obvious when seen through the ophthalmoscope, both of them
the consequence of what seems like a massive error of judgement on the part of Nature, namely the decision to have the retina inside out. The tips of the photoreceptors face outward, and they pass their information backwards through the retina: as a consequence, the nerve fibres from the retina find themselves inside the eye when they want to be outside. What they do is to come together to form the optic nerve, and crash their way to the exterior, together with the central retinal artery and vein, through a region called the optic disc, at about 15° to the nasal side of the optical axis. Since this area is consequently incapable of responding to light, subjectively it forms the blind spot. Although it is some 5° across, one is usually unaware of its existence because the brain tends to fill it in with whatever background colour or pattern immediately surrounds it. Close your left eye, and view the red cross in the figure below from a distance of about 30 cm: the alien will disappear, yet no discontinuity in the background will be apparent. image








Table 7.3 Factors affecting visual acuity



























































Target


Optical


Receptor/neural


Contrast


Aberrations


Receptor spacing: matched to best optics





Chromatic





Spherical


Colour


Diffraction


Neural convergence and divergence: decreased spatial resolution with dark adaptation



Wavelength – decreased diffraction if shorter; Monochromatic light – decreased chromatic aberration


Luminance


Refractive error



Very low luminance: increased quantum fluctuation



Myopia


Hypermetropia



Dark adaptation: large pupil degrades optics; increased neural convergence



Astigmatism




Glare, scatter




Pupil size





Decrease: increases diffraction





Increase: all other optical factors worse





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The other gross feature of the retina that is visible with the ophthalmoscope is that very close to the centre of the retina is an area about 15° across that is free of large blood vessels – they arch around on each side to supply it from the edge – and is also distinctly yellower than the rest of the field. This is the macula lutea (yellow spot), and at its centre is a very small dot – actually a depression or pit – called the fovea centralis. When we look at a small object in the outside world, it is the fovea that is directed to the corresponding part of the retinal image. This region is specialized for high-quality, photopic vision: the central
fovea is quite without rods, and the cones themselves are tightly packed to give the maximum information about image detail: the angular size of the rod-free region is about that of one’s fingernail with the hand fully extended. Cones in this region are about 2.3 µm across, corresponding to a visual angle of some half minute of arc. In the section of central monkey retina below, you can see that the depression arises because the retinal structures that elsewhere in the retina lie between the receptors and the lens – remembering again that the retina is inside-out in its layered structure – are here displaced to one side so as to cause the minimum scattering of incoming light. The supply of oxygen and nutrients for this region must derive almost entirely from the blood vessels that richly supply the choroid, the layer immediately superficial to the receptors and separated from them by the thin pigment epithelium, which helps to reduce scatter by absorbing the incident light. These features can all be seen in the section below, from a monkey retina.




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The retina is quite different from any of the sense organs we have met so far, in that a good deal of the neural processing of the afferent information has already occurred before it reaches the fibres of the optic nerve; it is in effect part of the brain. No doubt the reason for this is that the eye is a highly mobile organ, and if each of the 130 million or so receptors sent its own individual fibre into the optic nerve, the latter would have to be some 11 times thicker than at present, and would be a considerable hindrance to rapid movement of the eye; and of course the blind spot would be correspondingly larger as well. Thus some degree of compression of the afferent visual information is needed, and overall there is indeed a 100-fold convergence of information from large groups of receptors, particularly from rods in the periphery. The fibres of the optic nerve are in fact at two synapses’ remove from the cones: receptors synapse with bipolar cells, and these in turn synapse with the million or so ganglion cells whose axons form the optic nerve. These two types of neuron form consecutive layers on top of the receptor layer – except in the fovea, where we have seen that they are pushed to one side – and are mingled with two other types of interneuron that make predominantly sideways connections. These are the horizontal cells at the bipolar/receptor level, and, at the ganglion cell/bipolar level, the amacrine cells (some of which also provide another stage of convergence for the rod signals: see p. 145). The arrangement of the connections of all these types of interneuron is shown schematically above; the general arrangement is quite constant across species, though details vary. We shall see that there are marked differences in the electrical behaviour of these different neurons: although ganglion cells and amacrines show action potentials in response to retinal stimulation, the bipolars, horizontal cells and the receptors themselves do not. They are small enough to be able to interact electrotonically without the need for active propagation. image




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The photoreceptors

Rods and cones both consist of two distinct parts: an outer segment, apparently a grossly modified cilium, and an inner segment containing the nucleus. The outer segment possesses a high concentration of photopigment, associated with a richly folded set of invaginations of the outer surface, which are formed at the bottom and gradually move up to the tip over the course of a month
or so, then breaking off and being destroyed. In the case of rods they seal themselves off near the bottom, to form a stack of flattened saccules or discs; in the cones they remain partially open: it is not obvious why. At the base of the outer segment the remains of the ciliary filaments and centrioles can be seen. The inner segment has mitochondria as well as the nucleus, and its inner end forms the synaptic junction with bipolar and horizontal cells. There is no doubt that the photopigment straddling the membranes of the outer segment discs plays a key role in transforming incident light into electrical changes, for if the pigment is isolated from the receptor it is found that its absorption of light of different wavelengths corresponds closely with the spectral sensitivity of the receptors themselves.




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Retinal photopigment consists of two portions: a chromophore called retinal or retinene (a derivative of retinol, better known as vitamin A), in association with a protein/oligosaccharide complex with a molecular weight of around 40 000, called an opsin. It is slight differences in the composition of the opsin part that give rise to the different spectral sensitivities of rods and cones. In the case of the rods, the combination of rod opsin with retinal is called rhodopsin, sometimes also known as visual purple. In frog rods, there are about 1.5 million rhodopsin molecules in each disc, and about 1700 discs per rod. image Much more is known about rhodopsin than other pigments, but there is no reason to believe they are essentially different. Rhodopsin absorbs over nearly all the visible spectrum, peaking in the green region, at around 500 nm. We can compare this with the spectral sensitivity of vision itself, by measuring the absolute threshold for lights of different wavelength, in the dark-adapted state when only the rods are operating. We saw earlier (p. 129) that this curve, the scotopic sensitivity curve, corresponds very closely to the absorption spectrum for rhodopsin, implying that absorption by the pigment is indeed the first step in the transduction process.

The first effect of light on rhodopsin is to cause an isomerism of the retinal from the normal, bent, 11-cis form to the straightened all-trans configuration. This in turn leads to a series of changes in the configuration of the rhodopsin, producing a number of more-or-less short-lived intermediates, at the end of which opsin and retinal part company, and the rhodopsin is said to be bleached. In the test-tube, this is the end of the matter; but in the retina, the ingredients are all recycled by enzymes present in the receptors and in the pigment epithelium that lies behind them. The first stage of this process consists of the reconversion of the free all-trans retinal back to the 11-cis form, a relatively slow process. The significance of these wanderings of pigment back and forth between receptor and pigment epithelium is unclear. image




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We shall see later that it is this slow regeneration of pigment that determines the long time-course of recovery of rod sensitivity during dark adaptation that has already been mentioned (p. 128). In bright light, most of the rhodopsin is in the bleached form: an equilibrium is reached in which the rate of bleaching equals the rate
of regeneration. Estimates of the amount of pigment in the receptors of a living eye during particular stimulus conditions may be made by the technique of retinal reflection densitometry, in which one measures the amount and spectral composition of the light scattered back from the retina when a light is shone into the eye. In this way it is possible to track continuously the amount of rod or cone pigment in bleached form under relatively natural visual conditions. Alternatively, in microdensitometry, the spectral absorptions of individual receptors may be measured in a preparation on a microscope slide. As far as we know, the reactions that occur in rods and cones are fundamentally similar, though the regeneration of cone pigment is substantially quicker than in rods, so that under photopic conditions a smaller fraction of the cone pigment is in the bleached state than is the case for rods: this is one of the reasons why the cones are able to function at much higher light levels.


Electrical responses to light

The nature of the basic transduction process was outlined in Chapter 3 (p. 50). One of the stages in the sequence of photopigment bleaching – it is not certain which – is coupled by a G-protein called transducin to the activation of phosphodiesterase (PDE), that converts cyclic guanosine monophosphate (cGMP) to GMP. Since cGMP tonically promotes the opening of sodium channels in the plasma membrane, the effect of light on the outer segment is to reduce sodium permeability by reducing the level of cGMP. Because the effect of light is to reduce the amount of cGMP, you can see that – perhaps paradoxically – light will cause a reduction in sodium permeability instead of an increase. As a result, the receptor hyperpolarizes from a resting value of some -30 mV to a maximum of -60 mV, when the response saturates because all the channels are closed. As in many such cascades, there is a huge amplification of effects along the way: in rods, each quantum absorbed appears to cause the breakdown of about a million cGMP molecules, although the next stage is a bit of an anticlimax, since it takes three cGMPs to open a channel. Measurements of the absolute threshold for seeing dim flashes of light when the eye is fully dark-adapted show that a single rod is capable of responding to a single absorbed photon. Individually, cones are an order of magnitude less sensitive (photopic vision as a whole is several orders of magnitude less sensitive, because it enjoys less convergence and pooling of neural signals). image




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A disadvantage of cascades is that they tend to be slow, and the time-course of the hyperpolarization generated by a brief flash of light is very prolonged – a characteristic of indirect transduction with a lengthy cascade – and shows a pronounced plateau with very large stimuli, corresponding to closure of all the sodium channels, as can be seen in the cone hyperpolarizations below, in response to flashes of intensities ranging from 0 to 4.2 log units.




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As a result, if one plots the size of this receptor potential as a function of the intensity of the flash one finds a characteristic S-shaped, or saturating, relationship. The effect of different levels of light adaptation is to shift this curve along the intensity axis, over nearly 3.5 log units, providing one of the mechanisms by which the sensitivity of the retina is adjusted to suit the prevailing luminance. Bright backgrounds shorten the response as well as reducing its size; the significance and mechanism of this are discussed later, (p. 147). image

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Sep 8, 2016 | Posted by in NEUROLOGY | Comments Off on Vision

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